Interrupting Insects

A reposting of an original article from The Scorpion and the Frog.

What do you think of when I say “communicate”? Most likely, you are imagining people communicating by an auditory mode (talking and listening, making expressive sounds) or by a visual mode (observing body language, reading and writing). As a species, humans inherently rely heavily on our hearing and vision to perceive the world around us and so it makes sense that we communicate with one another using these modalities. But animal species are incredibly diverse in their means of perceiving their worlds and their modes of communication. Because we have been so focused on studying animal signals that we can perceive, we have only recently begun to more actively explore animal communication in these other modes. One of these modes is soundless surface vibrations.

The photo is of an adult Tylopelta gibbera on a host plant stem
(photo (c) Rex Cocroft).

Despite the fact that we do not perceive most animal surface vibration signals around us, vibrational communication is very common, especially among insects and spiders. Rex Cocroft at the University of Missouri at Columbia and Rafa Rodríguez at the University of Wisconsin at Milwaukee point out in a review of vibrational communication that over 195,000 species of insects communicate using soundless surface vibrations. We can experience many of these substrate vibration signals by broadcasting them through a speaker as an airborne vibration (which we perceive as sound).

Vibrational signals serve a number of functions in the insect worlds. Social insects, like ants, termites, and bees, often use vibrational signals to coordinate foraging. Groups of juvenile thornbug treehoppers vibrate when a predator approaches, calling in the mother to defend them. Males of many species have been found to use vibrational signals to attract females and the females often use these signals to choose a mate.

Vibrational signals are carried through a solid substrate, so they can only travel as far as the substrate is continuous and they are affected by attributes of the substrate (like changes in density). Because of these constraints, most vibrational signals can only travel about the length of a human arm. Many insects that use vibrational communication live on host plants, and it is these host plants that transmit the vibration signals. These animals face many challenges in transmitting their signals to the intended recipient. For example, wind, rain, and environmental sounds can create competing vibrations (background noise). In addition to environmental background noise, the vibrational soundscape of a given plant stem will likely include many signaling individuals, often of many species. Not only are there difficulties in getting your signal to your intended audience, but there are also risks of eavesdropping predators and competitors.

Frédéric Legendre, Peter Marting and Rex Cocroft at the University of Missouri at Columbia, demonstrate the social complexities of vibrational communication in a new study of competitive signaling in a treehopper species, Tylopelta gibbera. Tylopelta gibbera is a small treehopper in the southern United States, Mexico and Guatemala, that only lives on plants from the Desmodium genus. Males will attract and court females with vibrational signals and interested females will respond to the male with vibrational signals of their own. However, many individuals can often be found on a single plant and if two signaling males are present, the receptive female will typically respond to both of them and only mate with one (generally the first one she encounters). What is a competing male to do?

Listen to a male Tylopelta gibbera advertisement signal here.

The researchers performed a series of experiments, in which they observed treehoppers on potted host plants in the lab. With this set-up, they could control the environmental conditions, decide the number of males and females on the plant, record vibrational signals and play them back. They found that once a male signals and detects a female response, he will actively search for her along the plant, alternating signals and steps in a “Marco Polo” mating game until he finds her. Males found the females almost twice as fast if they were the only male on the plant, indicating that the presence of a second male on the plant somehow interferes with their ability to locate the female. Also, when two males were on the plant, they produced a new signal type that was never produced by a lone male on a plant. Males that had no male competition only produced signals that had a whine sound, followed by a series of pulses (and the female would then immediately respond with a harmonic sound of her own). This male signal is called the advertisement signal. Males that had a competing male on the plant would produce an additional signal that was a short tonal note. Interestingly, these males often produced this second signal at the same time that their competitor was advertising himself. Hmmm… could this be a masking signal used to interrupt the competitor? How could you figure that out?

This figure from Legendre, Marting and Cocroft's 2012 Animal Behaviour paper shows
the whine and pulses of a male advertisement signal (top) and a histogram of when the
masking signal occurs in relation to the timing of the advertisement signal (bottom).

First, the researchers asked, “When do males produce this second signal?” The researchers put two males on a plant with one female and recorded their vibrations. They found that in this situation, males typically produced this second signal while his competitor was just beginning the pulse section of his advertisement signal. Next, the researchers played back recordings of male advertisement signals followed by female responses to a lone male on a plant. All of the males tested produced the masking signal during the pulse section of the male advertisement signal on the recording.

Don't you hate it when someone does this?

Next, the researchers asked, “How do females respond to this second signal?” On plants with one female and two males, females didn’t respond as much to advertisement signals overlapped by a second signal as they did to advertisement signals alone. The researchers then played recordings of male advertisement signals to lone females on the plants. Females responded significantly more often if the advertisement signal was not overlapped by a masking signal.

So, male treehoppers get an edge up on getting the girl by interrupting the other competing males. Sneaky buggers!

Want to know more? Check these out:

1. COCROFT, R., & RODRÍGUEZ, R. (2005). The Behavioral Ecology of Insect Vibrational Communication BioScience, 55 (4) DOI: 10.1641/0006-3568(2005)055[0323:TBEOIV]2.0.CO;2

2. Legendre, F., Marting, P., & Cocroft, R. (2012). Competitive masking of vibrational signals during mate searching in a treehopper Animal Behaviour, 83 (2), 361-368 DOI: 10.1016/j.anbehav.2011.11.003

3. A Japanese research team has harnessed this phenomenon to create a remote-control that makes annoying people stop talking. Find out more at the blog Gaines on Brains!

Can You Feel the Love Tonight? (A Guest Post)

A reposting of an article by Maggie Nannenhorn from March 14, 2016.

If you’re like me, you never truly realize how quiet winter is until all the sounds of spring come back in a chorus of celebration. Between the birds, crickets, and frogs, you can really hear the love in the air. So you can hear the love, but can you feel the love?

Wood frogs are known for their chorus of calls that sound like a duck laughing. Seriously, tell a duck a good knock-knock joke and that is what a male wood frog sounds like when trying to attract a mate. He makes the call by expanding his two vocal sacs, membranes of skin underneath the neck, forming a bubble-like appearance. When a female surfaces, drawn to the call, the male frog clasps onto her, causing her to lay her eggs. The male frog then externally fertilizes the eggs. This form of mating is termed amplexus. The use of the call in the reproduction ritual is well studied. However, it is possible the small ripple formed in the water from the expanding vocal sack is relaying information that influences the mating behavior of these frogs.

Male wood frog resting on the water surface. Image by Maggie Nannenhorn.

Male wood frog calling with vocal sac expanded.
Notice the ripple it creates in the water. Image by Maggie Nannenhorn.

In 2010, Gerlinde Höbel and Robb Kolodziej from the University of Wisconsin-Milwaukee conducted an experiment that explored the use of water surface waves in wood frog reproductive behavior. They hypothesized male wood frogs use ripples in the water to find female wood frogs to mate with, while female wood frogs use ripples in the water as indicators of harassing males.

Video of a wood frog calling by Maggie Nannenhorn.

Wood frogs have a very short mating period: only 1 to 3 days per year! This study occurred on April 1st - 2nd, which corresponded with the wood frogs’ natural mating period. The first component of the study was the observation of a pond containing more than 500 wood frogs in amplexus. Amplexus was determined by the presence of males clasping on to the backs of female frogs in the water. They learned males approach surface waves on the water and clasp onto the frog that caused the ripple. However, females move away from surface waves on the water and dive downward.

After preliminary observations, they developed an experiment to cause rippling of the water. The first experiment tested the effect of stimulation (dipping a wooden probe into the water) near male wood frogs. The males tested were randomly assigned to either a control group or an experimental group. The 34 males in the control group were simply observed, and the direction and pattern of movement was recorded. For the experimental group, a long wooden probe was dipped in and out of the water 25 cm away from a male frog for 10 seconds. The resulting ripple was meant to mimic a ripple caused by a female frog moving in the water. Based on the hypothesis, the male wood frogs should approach the ripple hoping to find a female to mate with. Of the 60 males in the experimental group, half were stimulated from the right and half were stimulated from the left. A circle diagram (depicted below) was used to map the direction the males moved.

Video of a wood frog approaching ripples by Gerlinde Höbel.

This figure shows: a) the control group and b) the experimental group.
A circle diagram representing the reproductively driven movement direction
of wood frogs (Lithobates sylvaticus) in a laboratory pool as a result of
stimulated surface waves on both the left and right sides.
Figure from: Höbel, G., & Kolodziej, R. C. (2013). Behaviour, 150(5), 471-483.

The females are difficult to observe in the field since they prefer to stay beneath the surface. So, the researchers set up a tank to test 4 breeding pairs of wood frogs. They tested the females both while in amplexus and while alone. They dipped wooden probes into the water to stimulate the females on both the left and the right side in turn. Their positions and directions were also recorded using a circle diagram.

So, what did they find? It turns out, their predictions were correct! The males would approach the ripple caused by the probing. This is likely because the ripple may indicate a competing male they want to drive away or a female they want to mate with. The females moved away from the ripples by either swimming away or diving underneath the water surface. This may reduce the amount of harassment they receive from males. If a female becomes the center of attention for too many males, she may drown from the weight of them all attempting to grab her. Besides, if a male is fit, he will likely be able to catch up to her and successfully mate with her despite her swimming away.

The mating calls and movement of the wood frogs affect the surface waves, and these waves are used to make sexual behavior choices. This spring, the chorus of love will still ring out through the reeds, and I encourage you to take a moment to stop and listen. When you’re stopped, take a moment to notice the waves of love bringing these wood frogs together. Hopefully this spring, we will all be feeling the love.



Reference:

Höbel, G., & Kolodziej, R. (2013). Wood frogs (Lithobates sylvaticus) use water surface waves in their reproductive behaviour Behaviour, 1-13 DOI: 10.1163/1568539X-00003062

Can You Feel the Love Tonight? (A Guest Post)

By Maggie Nannenhorn

If you’re like me, you never truly realize how quiet winter is until all the sounds of spring come back in a chorus of celebration. Between the birds, crickets, and frogs, you can really hear the love in the air. So you can hear the love, but can you feel the love?

Wood frogs are known for their chorus of calls that sound like a duck laughing. Seriously, tell a duck a good knock-knock joke and that is what a male wood frog sounds like when trying to attract a mate. He makes the call by expanding his two vocal sacs, membranes of skin underneath the neck, forming a bubble-like appearance. When a female surfaces, drawn to the call, the male frog clasps onto her, causing her to lay her eggs. The male frog then externally fertilizes the eggs. This form of mating is termed amplexus. The use of the call in the reproduction ritual is well studied. However, it is possible the small ripple formed in the water from the expanding vocal sack is relaying information that influences the mating behavior of these frogs.

Male wood frog resting on the water surface. Image by Maggie Nannenhorn.

Male wood frog calling with vocal sac expanded.
Notice the ripple it creates in the water. Image by Maggie Nannenhorn.

In 2010, Gerlinde Höbel and Robb Kolodziej from the University of Wisconsin-Milwaukee conducted an experiment that explored the use of water surface waves in wood frog reproductive behavior. They hypothesized male wood frogs use ripples in the water to find female wood frogs to mate with, while female wood frogs use ripples in the water as indicators of harassing males.

Video of a wood frog calling by Maggie Nannenhorn.

Wood frogs have a very short mating period: only 1 to 3 days per year! This study occurred on April 1st - 2nd, which corresponded with the wood frogs’ natural mating period. The first component of the study was the observation of a pond containing more than 500 wood frogs in amplexus. Amplexus was determined by the presence of males clasping on to the backs of female frogs in the water. They learned males approach surface waves on the water and clasp onto the frog that caused the ripple. However, females move away from surface waves on the water and dive downward.

After preliminary observations, they developed an experiment to cause rippling of the water. The first experiment tested the effect of stimulation (dipping a wooden probe into the water) near male wood frogs. The males tested were randomly assigned to either a control group or an experimental group. The 34 males in the control group were simply observed, and the direction and pattern of movement was recorded. For the experimental group, a long wooden probe was dipped in and out of the water 25 cm away from a male frog for 10 seconds. The resulting ripple was meant to mimic a ripple caused by a female frog moving in the water. Based on the hypothesis, the male wood frogs should approach the ripple hoping to find a female to mate with. Of the 60 males in the experimental group, half were stimulated from the right and half were stimulated from the left. A circle diagram (depicted below) was used to map the direction the males moved.

Video of a wood frog approaching ripples by Gerlinde Höbel.

This figure shows: a) the control group and b) the experimental group.
A circle diagram representing the reproductively driven movement direction
of wood frogs (Lithobates sylvaticus) in a laboratory pool as a result of
stimulated surface waves on both the left and right sides.
Figure from: Höbel, G., & Kolodziej, R. C. (2013). Behaviour, 150(5), 471-483.

The females are difficult to observe in the field since they prefer to stay beneath the surface. So, the researchers set up a tank to test 4 breeding pairs of wood frogs. They tested the females both while in amplexus and while alone. They dipped wooden probes into the water to stimulate the females on both the left and the right side in turn. Their positions and directions were also recorded using a circle diagram.

So, what did they find? It turns out, their predictions were correct! The males would approach the ripple caused by the probing. This is likely because the ripple may indicate a competing male they want to drive away or a female they want to mate with. The females moved away from the ripples by either swimming away or diving underneath the water surface. This may reduce the amount of harassment they receive from males. If a female becomes the center of attention for too many males, she may drown from the weight of them all attempting to grab her. Besides, if a male is fit, he will likely be able to catch up to her and successfully mate with her despite her swimming away.

The mating calls and movement of the wood frogs affect the surface waves, and these waves are used to make sexual behavior choices. This spring, the chorus of love will still ring out through the reeds, and I encourage you to take a moment to stop and listen. When you’re stopped, take a moment to notice the waves of love bringing these wood frogs together. Hopefully this spring, we will all be feeling the love.



Reference:

Höbel, G., & Kolodziej, R. (2013). Wood frogs (Lithobates sylvaticus) use water surface waves in their reproductive behaviour Behaviour, 1-13 DOI: 10.1163/1568539X-00003062

Who Said What? (A Guest Post)

By Porscha Carriveau




A Quaker parrot shows off his beak
and tongue. Photo by Alex Nelson
at Wikimedia Commons.

As an aviculturist-turned-scientist, to me, it is common sense to tell people that birds are heard more often than seen. People study bird songs or calls for a variety of reasons. The reason I study bird songs is to identify the songs that my African grey parrot has learned to mimic. His repertoire includes the vocalizations of several birds’ songs such as robins, cardinals, cat birds, and chickadees. He also mimics humans. When leaving home in the morning, the last thing that I hear heading out the door is "gotta go to work" and the sound of being blown a kiss. Most people would think nothing of it, but I am being told this by a bird that has no lips.

Here is an example of an African grey parrot producing sound :

Humans produce sound by using their vocal tract, which includes the larynx (known as the voice box), where the vocal folds are located. Sound is produced with the help of the trachea, which controls air flow through the larynx. In the larynx the vocal folds make sound by vibrating. The remainder of the vocal tract includes the throat, nose, tongue and lips which are involved in the articulation of speech. On the other hand, parrots have a syrinx (what rivals the larynx), a trachea, a tongue and a beak. This means that birds do not have vocal cords to produce the sounds that we as humans make; they instead have two air passages that come together at the organ known as the syrinx creating a vibration that produces sound.

From my experiences working with and owning a variety of parrots, I would say that African grey parrots and monk parakeets (also known as Quaker parrots) are the two clearest and best mimicking parrots. Quaker parrots originate from South America. Over the years these birds have learned to adapt to their environment extremely well, leading to the birds becoming an invasive species in many parts of the world, including several U.S. states where they are now illegal to own as pets.

Research done by Verena Ohms, Gabriël Beckers, Carel ten Cate and Roderick Suthers recently set up a study using x-ray imaging to determine what is taking place in the vocal tract of a Quaker parrot while producing species specific calls. To do this, a piece of metal wire was placed on the underside of a Quaker parrot’s tongue and two pieces of wire were placed inside the trachea attached to tracheal rings. Here is an example of what researchers were looking at which allowed them to monitor the bird’s tongue, beak, and trachea movements.

Researchers looked specifically at a few measures when a bird produces sound: the bird’s tongue height (TH), the size of the beak opening (BO), and the amount of tracheal stretching (TS).

Diagram of the measures taken from Quaker parrots. Figure from Ohms, et al., 2012.

Through observing the changes that occurred from the metal wires placed inside a Quaker parrot’s tongue and trachea while producing calls, researchers were able to conclude that a parrot's tongue functions much differently than a songbirds’. Even more amazing is that a parrot’s tongue is similar to a human tongue in the way that it is manipulated while producing sound. Researchers also determined that these parrots manipulate the sound frequency (pitch) of their calls by moving their tongues in and out. The researchers were also the first to observe a circle-like movement in the trachea that had not been described before in this species.

So whether my trouble-making parrot (you should hear him burp and excuse himself) is blowing me a kiss or mimicking a bird song, there are many similarities in the way that humans and parrots produce speech sounds. This is pretty amazing for two groups of animals that are so different!


Work Cited

Ohms, V., Beckers, G., Ten Cate, C., & Suthers, R. (2012). Vocal Tract Articulation Revisited: The Case of the Monk Parakeet The Journal of Experimental Biology, 215, 85-92 DOI: 10.1242/jeb.064717

Red-Eyed Rump Shaker

A photo of a red-eyed treefrog taken
by Carey James Balboa at Wikimedia.

At night, male red-eyed treefrogs gather on saplings over Central American forest ponds to show off their stuff for the ladies, producing self-advertising “chack” calls. Despite the fact that they gather in groups, they defend their calling territories from flirtatious male competition. Females assess the available males and usually mate with a single male, who mounts her and clings on for dear life in a behavior called amplexus until she lays eggs that he then fertilizes. Occasionally, multiple males will try to mate with the same female at the same time, which usually results in two ticked-off male frogs.

What does an angry red-eyed treefrog do, you may ask? A mildly irritated treefrog will likely produce territorial “chuckle” calls, to let rivals know this is his favorite calling-plant and they’d better step-off. But a really ticked-off red-eyed treefrog rapidly lifts and lowers his hind end in a behavior called tremulation.

 
"Step off, I'm doing the hump!" Video by Michael Caldwell.

 Not much is known about this tremulation behavior. Is it something they do just to release anxiety or is it a communication signal? If it is a communication signal, is it a visual signal or a vibrational signal or both? And what exactly might it communicate?

Michael Caldwell, Karen Warkentin and Gregory McDaniel from Boston University, and Gregory Johnston from Flinders University in Australia, set out to ask the red-eyed treefrog if the tremulations were a communication signal and what they may mean. But without Dr. Doolittle’s powers of talking to the animals, how can scientists determine what and how animals are communicating?

First, the researchers observed natural interactions between males at choruses in the wild and recorded everything they did. They found that male red-eyed treefrogs will often approach another male while making “chuckles” and “chacks”. These males also tremulated in every aggressive interaction observed. Some of these males kicked with their back legs and some encounters even escalated to wrestling. Eventually (usually anywhere from a minute to an hour later, but occasionally several hours later), one of the males would submit by fleeing the plant or remaining silent and motionless. The dominant male would then resume his self-advertising “chack” calls. So males use tremulation in aggressive contexts with other males, but does that mean that it is a signal?

Males that won encounters tremulated more and used more
"chack" and "chuckle" calls than did males that lost encounters.
Figure from Caldwell et al. 2010 Current Biology paper.

The researchers then conducted staged contests by placing pairs of calling males on the same sapling. In these staged contests, males showed all the same aggressive behaviors the researchers had observed in natural conditions, and most ended in a wrestling match. The males that won their encounter produced more calls and more tremulations than did males that lost their encounter (Check out the graph above). So tremulations are used in the context of aggression with other males and winners tremulate more than losers. It looks like these tremulations are an aggressive communication signal, but to know for sure, we need to know if other males respond to them. And are tremulations a visual signal, a vibrational signal, or both?

So the researchers had to get creative and take it one step further: They put a robotic frog on a vibrating shaker that could mimic the visual display of a tremulation. They attached a separate vibrating shaker to the plant to mimic the vibrations of a tremulation. Now, they could look at the effects of the visual and vibrational components of the tremulation behavior separately!



Robofrog! Notice the jointed limbs and the metal rod sticking out of the robot's
belly. That rod is connected to a shaker that moves the robot so it looks like
he is performing a tremulation display. A separate shaker is connected to the
sapling to send the vibrational component of the display. This way, the
researchers can expose frogs to the visual component and the vibrational
component of the tremulation display separately. Photo by Michael Caldwell.

The researchers compared male red-eyed treefrogs that were exposed to (1) nothing, (2) a frog robot that does nothing, (3) a frog robot that “tremulates” with both plant vibrations and visible movement, (4) “tremulation” vibrations in the plant, without the frog robot, (5) a frog robot that moves it’s butt up and down but doesn’t produce vibrations, and (6) white noise vibrations in the plant (this is just a generic vibration).

Males responded aggressively to the imitated tremulation vibrations, visual or combined but not to any of the other treatments. This suggests that tremulations are a communication signal that rival males respond to. Interestingly, males only tremulated in response to tremulation vibrations. This suggests that the vibrational component is important to sending the full aggressive signal.

Males have aggressive responses to the visual display alone,
the vibration alone, and the visual display combined with the
vibration. But males only tremulated in response to vibrational
signals. Figure from Caldwell et al. 2010 Current Biology paper.

The sensitivity to soundless surface vibrations is widespread among animals, but we know very little about vibrational communication, especially in vertebrates. Michael, Gregory, Gregory and Karen have cleverly shown us that male red-eyed treefrogs use vibrational signals in contests with each other. How many other species will we discover using this silent channel of communication if we just listen?

Want to know more? Check this out:

Caldwell MS, Johnston GR, McDaniel JG, & Warkentin KM (2010). Vibrational signaling in the agonistic interactions of red-eyed treefrogs. Current biology : CB, 20 (11), 1012-7 PMID: 20493702

Interrupting Insects

What do you think of when I say “communicate”? Most likely, you are imagining people communicating by an auditory mode (talking and listening, making expressive sounds) or by a visual mode (observing body language, reading and writing). As a species, humans inherently rely heavily on our hearing and vision to perceive the world around us and so it makes sense that we communicate with one another using these modalities. But animal species are incredibly diverse in their means of perceiving their worlds and their modes of communication. Because we have been so focused on studying animal signals that we can perceive, we have only recently begun to more actively explore animal communication in these other modes. One of these modes is soundless surface vibrations.

The photo is of an adult Tylopelta gibbera on a host plant stem
(photo (c) Rex Cocroft).

Despite the fact that we do not perceive most animal surface vibration signals around us, vibrational communication is very common, especially among insects and spiders. Rex Cocroft at the University of Missouri at Columbia and Rafa Rodríguez at the University of Wisconsin at Milwaukee point out in a review of vibrational communication that over 195,000 species of insects communicate using soundless surface vibrations. We can experience many of these substrate vibration signals by broadcasting them through a speaker as an airborne vibration (which we perceive as sound).

Vibrational signals serve a number of functions in the insect worlds. Social insects, like ants, termites, and bees, often use vibrational signals to coordinate foraging. Groups of juvenile thornbug treehoppers vibrate when a predator approaches, calling in the mother to defend them. Males of many species have been found to use vibrational signals to attract females and the females often use these signals to choose a mate.

Vibrational signals are carried through a solid substrate, so they can only travel as far as the substrate is continuous and they are affected by attributes of the substrate (like changes in density). Because of these constraints, most vibrational signals can only travel about the length of a human arm. Many insects that use vibrational communication live on host plants, and it is these host plants that transmit the vibration signals. These animals face many challenges in transmitting their signals to the intended recipient. For example, wind, rain, and environmental sounds can create competing vibrations (background noise). In addition to environmental background noise, the vibrational soundscape of a given plant stem will likely include many signaling individuals, often of many species. Not only are there difficulties in getting your signal to your intended audience, but there are also risks of eavesdropping predators and competitors.

Frédéric Legendre, Peter Marting and Rex Cocroft at the University of Missouri at Columbia, demonstrate the social complexities of vibrational communication in a new study of competitive signaling in a treehopper species, Tylopelta gibbera. Tylopelta gibbera is a small treehopper in the southern United States, Mexico and Guatemala, that only lives on plants from the Desmodium genus. Males will attract and court females with vibrational signals and interested females will respond to the male with vibrational signals of their own. However, many individuals can often be found on a single plant and if two signaling males are present, the receptive female will typically respond to both of them and only mate with one (generally the first one she encounters). What is a competing male to do?

Listen to a male Tylopelta gibbera advertisement signal here.

The researchers performed a series of experiments, in which they observed treehoppers on potted host plants in the lab. With this set-up, they could control the environmental conditions, decide the number of males and females on the plant, record vibrational signals and play them back. They found that once a male signals and detects a female response, he will actively search for her along the plant, alternating signals and steps in a “Marco Polo” mating game until he finds her. Males found the females almost twice as fast if they were the only male on the plant, indicating that the presence of a second male on the plant somehow interferes with their ability to locate the female. Also, when two males were on the plant, they produced a new signal type that was never produced by a lone male on a plant. Males that had no male competition only produced signals that had a whine sound, followed by a series of pulses (and the female would then immediately respond with a harmonic sound of her own). This male signal is called the advertisement signal. Males that had a competing male on the plant would produce an additional signal that was a short tonal note. Interestingly, these males often produced this second signal at the same time that their competitor was advertising himself. Hmmm… could this be a masking signal used to interrupt the competitor? How could you figure that out?

This figure from Legendre, Marting and Cocroft's 2012 Animal Behaviour paper shows
the whine and pulses of a male advertisement signal (top) and a histogram of when the
masking signal occurs in relation to the timing of the advertisement signal (bottom).

First, the researchers asked, “When do males produce this second signal?” The researchers put two males on a plant with one female and recorded their vibrations. They found that in this situation, males typically produced this second signal while his competitor was just beginning the pulse section of his advertisement signal. Next, the researchers played back recordings of male advertisement signals followed by female responses to a lone male on a plant. All of the males tested produced the masking signal during the pulse section of the male advertisement signal on the recording.

Don't you hate it when someone does this?

Next, the researchers asked, “How do females respond to this second signal?” On plants with one female and two males, females didn’t respond as much to advertisement signals overlapped by a second signal as they did to advertisement signals alone. The researchers then played recordings of male advertisement signals to lone females on the plants. Females responded significantly more often if the advertisement signal was not overlapped by a masking signal.

So, male treehoppers get an edge up on getting the girl by interrupting the other competing males. Sneaky buggers!

Want to know more? Check these out:

1. COCROFT, R., & RODRÍGUEZ, R. (2005). The Behavioral Ecology of Insect Vibrational Communication BioScience, 55 (4) DOI: 10.1641/0006-3568(2005)055[0323:TBEOIV]2.0.CO;2

2. Legendre, F., Marting, P., & Cocroft, R. (2012). Competitive masking of vibrational signals during mate searching in a treehopper Animal Behaviour, 83 (2), 361-368 DOI: 10.1016/j.anbehav.2011.11.003

3. A Japanese research team has harnessed this phenomenon to create a remote-control that makes annoying people stop talking. Find out more at the blog Gaines on Brains!